Canopy-forming Algae Research Articles

Recent and historical range shifts of two canopy-forming seaweeds in North Spain and the link with trends in sea surface temperature

Geographical range shifts of two canopy-forming seaweeds, Himanthalia elongata (L.) S.F. Gray and Fucus serratus L. were investigated at their southern range boundary in Northern Spain from the end of nineteenth century to 2009. Given the good dispersal abilities of H. elongata and its short life-span, we hypothesize that this species will track environmental changes at a faster rate than the perennial and short-distance disperser F. serratus. Our results show a continuous and drastic westward retraction of H. elongata, which has nowadays virtually vanished in Northern Spain, whereas F. serratus is still found in the westernmost area. Despite this, the first species is still relatively abundant in the Iberian Peninsula, whereas the presence of the latter is scattered and reduced. Overall, range shifts fit with the warming trend in sea surface temperature (SST), whereas it is unlikely that increases in grazing pressure or coastal pollution have driven the observed changes, particularly the rapid trend in recent years. Differences in species traits are linked to range dynamics. The higher persistence of F. serratus at eastern shores may thus be related to its longer life span and its greater thermal tolerance. The presence of sporadic populations of H. elongata outside the zone of continuous distribution can be attributed to long-distance dispersal events during cold pulses. Relict populations in isolated and estuarine locations were left behind in the contracting range margins, particularly for F. serratus. In Northern Spain, the westward retreat of large canopy-forming algae seems to be a general phenomenon, involving other species such as kelps. Therefore, an evident reorganization of coastal assemblages is expected, though the temporal extent of changes and the consequences for ecosystem services need to be evaluated.

Habitat-Associations of Turban Snails on Intertidal and Subtidal Rocky Reefs

Patchiness of habitat has important influences on distributions and abundances of organisms. Given the increasing threat of loss and alteration of habitats due to pressures associated with humans, there is a need for ecologists to understand species' requirements for habitat and to predict changes to taxa under various future environmental conditions. This study tested hypotheses about the generality of patterns described for one species of marine intertidal turban snail for a different, yet closely-related species in subtidal habitats along the coast of New South Wales, Australia. These two closely-related species live in similar habitats, yet under quite different conditions, which provided an opportunity to investigate how similar types of habitats influence patterns of distribution, abundance and size-structure in intertidal versus subtidal environments. For each species, there were similar associations between biogenically structured habitat and densities. The intertidal species, Turbo undulates, were more abundant, with greater proportions of small individuals in habitats formed by the canopy-forming alga, Hormosira banksii, the solitary ascidian, Pyura stolonifera or the turfing red alga, Corallina officinalis compared to simple habitat (bare rock). Similarly, more Turbo torquatus were found in biogenically structured subtidal habitat, i.e. canopy-forming algae, Ecklonia radiata, mixed algal communities (‘fringe’), or turfing red algae (Corallina officinalis and Amphiroa aniceps) than where habitat is simple (barrens). Small T. torquatus were more abundant in areas of turf and ‘fringe’, while large snails were more abundant in areas of kelp and barrens. These patterns were found at each location sampled (i.e. eight intertidal and two subtidal rocky reefs) and at all times of sampling, across each environment. This study highlighted the consistent influence of biogenically structured habitats on the distribution, abundance and size-structure of intertidal and subtidal turban snails and forms a basis for increasing the understanding of the potential underlying processes causing such patterns.

Shift happens: trailing edge contraction associated with recent warming trends threatens a distinct genetic lineage in the marine macroalga Fucus vesiculosus

BackgroundSignificant effects of recent global climate change have already been observed in a variety of ecosystems, with evidence for shifts in species ranges, but rarely have such consequences been related to the changes in the species genetic pool. The stretch of Atlantic coast between North Africa and North Iberia is ideal for studying the relationship between species distribution and climate change as it includes the distributional limits of a considerable number of both cold- and warm-water species.We compared temporal changes in distribution of the canopy-forming alga Fucus vesiculosus with historical sea surface temperature (SST) patterns to draw links between range shifts and contemporary climate change. Moreover, we genetically characterized with microsatellite markers previously sampled extinct and extant populations in order to estimate resulting cryptic genetic erosion.ResultsOver the past 30 years, a geographic contraction of the southern range edge of this species has occurred, with a northward latitudinal shift of approximately 1,250 km. Additionally, a more restricted distributional decline was recorded in the Bay of Biscay. Coastal SST warming data over the last three decades revealed a significant increase in temperature along most of the studied coastline, averaging 0.214°C/decade. Importantly, the analysis of existing and extinct population samples clearly distinguished two genetically different groups, a northern and a southern clade. Because of the range contraction, the southern group is currently represented by very few extant populations. This southern edge range shift is thus causing the loss of a distinct component of the species genetic background.ConclusionsWe reveal a climate-correlated diversity loss below the species level, a process that could render the species more vulnerable to future environmental changes and affect its evolutionary potential. This is a remarkable case of genetic uniqueness of a vanishing cryptic genetic clade (southern clade).

Fine-scale effects of sedentary urchins on canopy and understory algae

The effect of sea urchins on algae has often been studied when sea urchins forage over areas to create or maintain urchin barrens. The effect of sea urchins that are not associated with barrens, however, is not so clear. This study tested the hypothesis that sedentary and largely site attached sea urchins (Heliocidaris erythrogramma) have detectable effects on the covers of canopy-forming and understory algae, whether that be due to their mere presence or grazing. Algal cover was estimated beneath and immediately surrounding urchins in a location where they do not create barrens but attain densities at the upper end of their local range. The area immediately beneath the urchins was mostly devoid of visible algae. Experimental removal of urchins over 3years revealed that canopy-forming algae were able to develop in more extensive covers to match those in areas naturally lacking urchins. In the presence of urchins, the immediate area (i.e. first 5–6cm from urchins) was primarily comprised of turf-forming algae, thereafter the cover of canopy-forming algae sharply increased until at ~8cm their cover matched areas without urchins. These data support the model that composition and cover of algae can be altered in the presence of non-barren forming urchins, albeit only within a few centimetres of individual urchins. Where populations of sedentary urchins are dense and widespread (e.g. sheltered coasts for H. erythrogramma) such fine-scale effects may repeat over large areas such that urchins do not need to be associated with barrens before their effects are detectable.

Variation in community structure across vertical intertidal stress gradients: how does it compare with horizontal variation at different scales?

In rocky intertidal habitats, the pronounced increase in environmental stress from low to high elevations greatly affects community structure, that is, the combined measure of species identity and their relative abundance. Recent studies have shown that ecological variation also occurs along the coastline at a variety of spatial scales. Little is known, however, on how vertical variation compares with horizontal variation measured at increasing spatial scales (in terms of sampling interval). Because broad-scale processes can generate geographical patterns in community structure, we tested the hypothesis that vertical ecological variation is higher than fine-scale horizontal variation but lower than broad-scale horizontal variation. To test this prediction, we compared the variation in community structure across intertidal elevations on rocky shores of Helgoland Island with independent estimates of horizontal variation measured at the scale of patches (quadrats separated by 10s of cm), sites (quadrats separated by a few m), and shores (quadrats separated by 100s to 1000s of m). The multivariate analyses done on community structure supported our prediction. Specifically, vertical variation was significantly higher than patch- and site-scale horizontal variation but lower than shore-scale horizontal variation. Similar patterns were found for the variation in abundance of foundation taxa such as Fucus spp. and Mastocarpus stellatus, suggesting that the effects of these canopy-forming algae, known to function as ecosystem engineers, may explain part of the observed variability in community structure. Our findings suggest that broad-scale processes affecting species performance increase ecological variability relative to the pervasive fine-scale patchiness already described for marine coasts and the well known variation caused by vertical stress gradients. Our results also indicate that experimental research aiming to understand community structure on marine shores should benefit from applying a multi-scale approach.

Recovery of canopy-forming macroalgae following removal of the enigmatic grazing sea urchin Heliocidaris erythrogramma

The sea urchin Heliocidaris erythrogramma (Valenciennes, 1846) is considered an ecologically important member of shallow sub-tidal reef assemblages across temperate Australia. However, defining its ecological role has remained elusive due to a paucity of evidence demonstrating the ability of this species to graze and maintain sea urchin barrens. Here we present critical evidence for a grazing effect of H. erythrogramma at typically observable densities (~ 4–6 individuals m − 2 ) by demonstrating recovery of canopy-forming brown algae following two controlled removals of the sea urchin from ostensibly barrens reef at sheltered locations in eastern Tasmania. In experimental plots where H. erythrogramma was removed, canopy-forming algae gradually recovered to demonstrate an average 5 times (ranging nil to 10 times) increase in percentage cover at 24 months post manipulation (chiefly driven by the habitat-formers Cystophora spp., Macrocystis pyrifera, Acrocarpia paniculata, and Sargassum spp.). While divergence in the overall algal community was indicated by both experiments, a statistically significant shift, based on percentage cover community data, was observed for only one location at 24 months post sea urchin removal, suggesting that a complete canopy-driven shift in community structure will be gradual and will be contingent on urchin density remaining below approximately 1.5 m − 2 in the longer term. The relatively slow and highly variable rates of algal recovery across plots suggests that seasonal coexistence of annual (chiefly filamentous algae) and some perennial macroalgae on H. erythrogramma barrens is likely due to a relatively low and variable intensity of selective grazing. We conclude that H. erythrogramma, at sufficient densities, is capable of grazing and maintaining a sea urchin barrens state however this phenomenon is observed to manifest in sheltered waters only. In contrast, H. erythrogramma occurring on exposed Tasmanian reefs achieves similar densities but here barrens are not observed and the sea urchin occurs cryptically within crevices and, consistent with experiments elsewhere in Australia, the sea urchin under these conditions appears to feed exclusively on an abundance of detached ‘drift’ macroalgae. Top–down and bottom–up factors leading to increased abundance of H. erythrogramma will be important in determining the potential influence of this enigmatic grazer on rocky reef communities, however critical tests of mechanisms controlling the switch between foraging modes are needed. Taken together, correlative patterns and results of experimental removals across the range of H. erythrogramma demonstrate that models of temperate Australian reef dynamics must account for the highly context-dependent effects of sea urchin foraging.

Loss and Recovery Potential of Marine Habitats: An Experimental Study of Factors Maintaining Resilience in Subtidal Algal Forests at the Adriatic Sea

BackgroundPredicting and abating the loss of natural habitats present a huge challenge in science, conservation and management. Algal forests are globally threatened by loss and severe recruitment failure, but our understanding of resilience in these systems and its potential disruption by anthropogenic factors lags well behind other habitats. We tested hypotheses regarding triggers for decline and recovery potential in subtidal forests of canopy-forming algae of the genus Cystoseira.Methodology/Principal FindingsBy using a combination of historical data, and quantitative in situ observations of natural recruitment patterns we suggest that recent declines of forests along the coasts of the north Adriatic Sea were triggered by increasing cumulative impacts of natural- and human-induced habitat instability along with several extreme storm events. Clearing and transplantation experiments subsequently demonstrated that at such advanced stages of ecosystem degradation, increased substratum stability would be essential but not sufficient to reverse the loss, and that for recovery to occur removal of the new dominant space occupiers (i.e., opportunistic species including turf algae and mussels) would be required. Lack of surrounding adult canopies did not seem to impair the potential for assisted recovery, suggesting that in these systems recovery could be actively enhanced even following severe depletions.Conclusions/SignificanceWe demonstrate that sudden habitat loss can be facilitated by long term changes in the biotic and abiotic conditions in the system, that erode the ability of natural ecosystems to absorb and recover from multiple stressors of natural and human origin. Moreover, we demonstrate that the mere restoration of environmental conditions preceding a loss, if possible, may be insufficient for ecosystem restoration, and is scarcely cost-effective. We conclude that the loss of complex marine habitats in human-dominated landscapes could be mitigated with appropriate consideration and management of incremental habitat changes and of attributes facilitating system recovery.

Land‐to‐sea connectivity: linking human‐derived terrestrial subsidies to subtidal habitat change on open rocky coasts

Spatial subsidies are considered strong where differences in resource availability between donor and recipient systems are greatest. We tested whether human activities on land can increase subsidies of terrigenous nitrogen to open rocky coasts and whether these differences can predict apparent deforestation of kelp forests. We first identified landscape-scale variation in the human-mediated transfer of dissolved inorganic nitrogen (DIN) from natural, agricultural, and urban catchments to temperate coasts after episodes of rainfall. Compared to natural catchments, subsidies of DIN were on average 8-407 times greater in urban catchments, and 1-63 times greater in agricultural catchments. Urban derived nitrogen was attributed to the release of sewage effluent, as delineated by delta15N isotopic values of transplanted algae. Having made this link, we then assessed whether this catchment-scale variation may account for variation in structure of subtidal habitats, particularly as related to theory of nutrient-driven shifts of habitat from perennial (i.e., canopy-forming algae) to opportunistic species (i.e., turf-forming algae). We not only detected patterns consistent with this theory, but also established that the size and total proportion of patches of turf-forming algae were greater where the ratio of donor: recipient nitrogen loads was greater (i.e., size of subsidy). An important realization was that deforestation may be more strongly related to variation in the size of subsidy rather than size of human populations, particularly among urban catchments. These data directly link the type of human activity within catchments to the modification of land-to-sea subsidies and indirectly support theory that predicts terrestrial inputs to have greater ecological effects where the disparity in resource availability between donor and recipient is exacerbated. Our evidence has been used by coastal managers to reconsider their management of coastal systems and has subsequently contributed to new water-recycling policy and initiatives.

Effects of mean intensity and temporal variability of disturbance on the invasion of Caulerpa racemosa var. cylindracea (Caulerpales) in rock pools

Disturbance is a key factor influencing the invasibility of habitats and assemblages. This relationship was extensively studied in terrestrial systems, but it was scarcely tested in the marine environment. We investigated experimentally the interactive effects of changes in the intensity and temporal variability of mechanical disturbance by boulders on invasion dynamics of the green alga Caulerpa racemosa var. cylindracea in littoral rock pools. We tested the hypothesis that the success of invasion of C. racemosa would be (1) greater under large than under low intensity of disturbance, (2) greater under large than under low temporal variability of disturbance and that (3) interactive effects could also occur, with variability of disturbance magnifying the effects of intensity. C. racemosa was virtually absent in pools maintained under high intensity of disturbance, independently of temporal variability. High intensity of disturbance was also associated with lower density and length of fronds and thinner diameter of the stolons of the alga. The total number of native taxa and the abundance of encrusting coralline algae increased under high intensity of disturbance. Differently, turf-forming algae were positively affected by temporal variability of disturbance, while canopy-forming algae did not respond to experimental treatments. Our results suggest a direct negative effect of the most severe experimental conditions on the spread of C. racemosa in rock pools. This likely overwhelmed likely concomitant positive and negative effects mediated by resident organisms. The results of this study help anticipating invasion dynamics of C. racemosa in rock pools under climate change scenarios, in which both intensity and temporal variability of extreme meteorological events are predicted to increase.

BENTHIC METABOLISM ON A SHELTERED ROCKY SHORE: ROLE OF THE CANOPY IN THE CARBON BUDGET(1).

While the importance of canopy-forming algae in structuring ecosystems is recognized, their role in the carbon budget is still not well understood. To our knowledge, no measurements of rocky shores primary production and respiration under emersion periods have been carried out in situ. A benthic chamber coupled to a CO2 -infrared gas analyzer was used to measure gross primary production and respiration on the Ascophyllum nodosum (L.) Le Jol. zone of a sheltered rocky shore in Brittany, France. Over a year of monthly measurements on the zone with and without the A.nodosum canopy showed fairly high production and respiration values for the global community as well as carbon fluxes due to the canopy that largely dominated the benthic metabolism of the zone. The strong canopy respiration relative to the primary production also suggested a high metabolic activity by microscopic heterotrophs on the surface of the alga. Both the canopy and the understory annual primary production and respiration were under the control of light and temperature seasonal variations. Finally, the range of the amount of carbon produced on the A.nodosum zone during diurnal emersions was estimated. Additional measures accounting for the day-night cycles and seasonal light variations over an entire tidal cycle are, however, necessary to establish an annual carbon budget. Such measures using the benthic chamber together with complementary techniques would allow a better understanding of the functioning of sheltered rocky shores.

Recovering a lost baseline: missing kelp forests from a metropolitan coast

There is concern about historical and continuing loss of canopy-forming algae across the world's temperate coastline. In South Australia, the sparse cover of canopy-forming algae on the Adelaide metropolitan coast has been of public concern with continuous years of anecdotal evidence culminating in 2 competing views. One view considers that current patterns existed before the onset of urbanisation, whereas the alternate view is that they developed after urbanisation. We tested hypothe- ses to distinguish between these 2 models, each centred on the reconstruction of historical covers of canopies on the metropolitan coast. Historically, the metropolitan sites were indistinguishable from con- temporary populations of reference sites across 70 km (i.e. Gulf St. Vincent), and could also represent a random subset of exposed coastal sites across 2100 km of the greater biogeographic province. Thus there was nothing 'special' about the metropolitan sites historically, but today they stand out because they have sparser covers of canopies compared to equivalent locations and times in the gulf and the greater province. This is evidence of wholesale loss of canopy-forming algae (up to 70%) on parts of the Adelaide metropolitan coast since major urbanisation. These findings not only set a research agenda based on the magnitude of loss, but they also bring into question the logic that smaller metropolitan populations of humans create impacts that are trivial relative to that of larger metropolitan centres. Instead, we highlight a need to recognise the ecological context that makes some coastal systems more vulnerable or resistant to increasing human-domination of the world's coastlines. We discuss challenges to this kind of research that receive little ecological discussion, particularly better leadership and administration, recognising that the systems we study out-live the life spans of individual research groups and operate on spatial scales that exceed the capacity of single research providers.

Physical disturbance and subtidal habitat structure on open rocky coasts: Effects of wave exposure, extent and intensity

Canopy-forming algae occur across of range of energy environments (i.e., wave sheltered to exposed coasts) where disturbances are frequent (i.e., gap formation) and benthic patterns largely reflect variation in post-disturbance processes. Disturbances vary in extent (area affected) and intensity (degree of damage), and this may affect recolonisation at local scales. On an open oceanic coast, we tested whether habitat structure (patches of canopy algae) differed between heavy and relatively lighter wave exposure (sheltered vs. exposed sides of islands), and whether wave exposure affected the response of prominent habitat-formers to varying disturbance regimes (different sizes of partial and complete canopy removal). Observations of naturally occurring patterns showed sheltered coasts to be characterised by small patches of fucoids, whereas exposed coasts were characterised by large patches of kelp. Canopy-gaps were larger at exposed than sheltered coasts, and mixed canopies constituted > 24% of the subtidal rocky habitat independently of wave exposure. Experimental disturbances showed the local density of kelps to affect recovery through greater recruitment to partial clearings (80% canopy removal). Fucalean algae, on the other hand, mainly recruited into complete clearings (100% removal), but when their recruits were abundant, they also recruited into partial clearings. The covers of filamentous, turf-forming algae increased in all clearings, and more so at exposed than sheltered sites. Extent of disturbance had no detectable effect on recolonisation by canopy-forming algae across the scales examined (i.e., 1.5 m, 3 m diameter loss of canopy). Recolonisation varied among islands kilometres apart, and correlations ( r > 0.85) between cover of canopies and cover of their recruits in clearings at the scale of sites, suggested that differences in propagule supply could account for variation in patterns of recolonisation at scales of kilometres. There was no evidence to suggest that the effect of disturbance depended on wave exposure within the range of exposures tested in this study (i.e. open coasts). We recognise that wave exposure can be fundamental to habitat structure of subtidal rocky coasts, but we suggest that its influence may be mediated by the biological setting (e.g., canopy composition).

The invasion of the intertidal canopy-forming alga Fucus serratus L. to southwestern Iceland: Possible community effects

The intertidal serrated wrack, Fucus serratus L. (Fucales: Phaeophyceae), has become an abundant canopy-forming alga along a ca. 100 km stretch of shore in southwestern Iceland in the last century. Its distribution has not changed noticeably since a survey in 1975/1976. Many experimental studies have shown that canopy-forming algae can have profound effects on the community structure. Although such experiments involving F. serratus are few it was decided to test predictions from these experiments on a geographical scale by comparing community compositions within the area where F. serratus is a dominant algae (since at least 1975/1976) with an adjacent area of similar size where F. serratus is absent (with a single exception without consequences). The work is based on measurements on some 372 stations in the F. serratus area, termed region A, and 227 stations from the F. serratus-free area, termed region B. Percentage cover of algae and sessile invertebrates was estimated on 2 m 2 on each station and animals collected from 800 cm 2 from each station. The vertical distribution of F. serratus was most similar to that of Fucus distichus and the two species often grew intermingled on the lower part of the shore. F. serratus appeared to have reduced the cover of F. distichus in the lowermost part of the shore, while having little or no effect on other canopy-forming species. The low abundance of Semibalanus balanoides in the F. serratus area (region A) is in line with experiments showing detrimental effect on this barnacle by whiplash of F. serratus. In general algal grazers were more abundant in region A, consistent with the greater attractiveness of F. serratus than other canopy-forming species to grazers, although this may be partly explained by the abundance of understorey algae in region A.

Mechanisms of recovery and resilience of different components of mosaics of habitats on shallow rocky reefs

Understanding the mechanisms producing and maintaining discontinuities between patches in mosaics of habitats is necessary to predict changes in patterns of abundance and distribution of species. On temperate rocky reefs, physical and biological disturbance can result in a mosaic of patches of encrusting coralline, turf- or canopy-forming algae. We experimentally investigated the effects of disturbance within and at the boundary between these patches, in order to identify mechanisms accounting for re-colonisation of space and to assess whether the response of boundaries can alter the relative size of contrasting habitats. Also, we tested whether the resilience of the different types of assemblages depends on species richness (i.e. number of taxa present) of habitats or, alternatively, on other properties of systems like differences in life-history traits of dominating species. Although the nature of the mechanisms generating differences among habitats changed among different stages of the colonisation, local processes (within patches) prevailed over larger scale processes (among patches) in determining early patterns of colonisation of space in mosaics. By the end of the experiment, assemblages in clearings at boundaries had recovered to the undisturbed reference condition, in contrast to clearings within barren patches or algal turfs. Boundaries represent, therefore, relatively more stable components of the mosaic, with greater resilience than adjacent habitats. Although sea urchins are often indicated as the main force regulating the proportion of contrasting habitats on shallow rocky reefs, determining the nature of variation in the interaction between species dominating each type of habitat is crucial to understand dynamics of mosaics. Finally, our study provides evidence that resilience could not be entirely controlled by initial species diversity, suggesting that different proportions of dominant taxa could influence the stability of natural systems.

Community effects following the deletion of a habitat-forming alga from rocky marine shores

Habitat-forming species increase spatial complexity and alter local environmental conditions, often facilitating a diversified assemblage of plants and animals. Removal of dominant species, therefore, can potentially lead to pronounced changes in diversity and community structure through a series of negative and positive interactions involving several components of the community. Here we test community responses to the deletion of the dominant, canopy-forming alga Hormosira banksii from the mid-intertidal zone of wave-protected rocky shores in southern New Zealand. This species was removed in winter (July) from three 3x3-m areas at each of two platforms (Kaikoura and Moeraki) on the east coast of the South Island. Initially, 59 taxa occurred in stands, but there were only four algal species with greater than 5% cover and three mobile invertebrate species with more than five individuals per 0.25 m(2). By 6 months after Hormosira removal, most fucoid and coralline algae had burned off, and there were blooms of ephemeral algae in the removal plots, but almost no change within controls. After 2 years, diversity declined by 44% relative to controls at Kaikoura and 36% at Moeraki, and the amount of bare space had increased by tenfold at Kaikoura and twofold at Moeraki. Few sessile or mobile invertebrates were present. Recruitment of Hormosira occurred after 14 months in the removal plots. At this time, a "press" disturbance was initiated into one half of each removal plot to test the effects of continued removal of Hormosira on diversity. Similar "end-points" of the control and "press" removal plots were not reached after 2 years, and even after Hormosira recruitment into the original "pulse" experiment there was little recovery of the community. In this mid-intertidal system with considerable thermal stress, and perhaps in others with few perennial species, diversity and community structure can critically depend on positive associations with a single dominant species.

Predicting understorey structure from the presence and composition of canopies: an assembly rule for marine algae.

Assembly rules provide a useful framework for predicting patterns of community assembly under defined environmental conditions. Habitat created by canopy-forming algae (such as kelps) provides a promising system for identifying assembly rules because canopies typically have a large and predictable influence on understorey communities. Across >1,000 km of subtidal South Australian coastline, we identified natural associations between assemblages of understorey algae and (1) monospecific canopies of Ecklonia radiata, (2) canopies comprised of E. radiata mixed with Fucales (Cystophora spp. and Sargassum spp.), and (3) gaps among canopies of algae. We were able to recreate these associations with experimental tests that quantified the assembly of understorey algae among these three habitat types. We propose the assembly rule that understorey communities on subtidal rocky coast in South Australia will be (1) monopolised by encrusting coralline algae beneath monospecific canopies of E. radiata, (2) comprised of encrusting corallines, encrusting non-corallines, and sparse covers of articulated corallines, beneath mixed E. radiata-Fucales canopies, and (3) comprised of extensive covers of articulated corallines and filamentous turfs, as well as sparse covers of foliose algae and juvenile canopy-formers, within gaps. Consistencies between natural patterns and experimental effects demonstrate how algal canopies can act as a filter to limit the subsets of species from the locally available pool that are able to assemble beneath them. Moreover, the subsets of species that assemble to subtidal rocky substrata in South Australia appear to be predictable, given knowledge of the presence and composition of canopies incorporating E. radiata.

Disturbance initiates diversity in recruitment of canopy-forming algae: interactive effects of canopy-thinning and substratum availability

P.J. Goodsell and S.D. Connell. 2005. Disturbance initiates diversity in recruitment of canopy-forming algae: interactive effects of canopy-thinning and substratum availability. Phycologia 44: 632–639.We assessed whether disturbance within monospecific canopies (Ecklonia radiata, Phaeophyta) increases diversity of macroalgal recruits by creating gaps in the canopy and by liberating space on the substratum. We tested the hypothesis that a greater number of species of canopy-forming algae recruit to areas where canopies of monospecific E. radiata are thinned and the availability of barer substratum is increased (i.e. fewer holdfasts without thallus remain attached to the substratum). A significant interaction between canopy-cover and substrate-availability was detected; losses to canopy caused greater diversity of recruits only where there were few holdfasts remaining on the substratum. Interestingly, we found large numbers of species under thinned canopies where remnant holdfasts were sparse, which were also found under dense canopies where remnant holdfasts were dense. Residual holdfasts may increase the likelihood of survival of recruits under dense canopies through the provision of habitat rather than displacing appropriate habitat for recruitment. Nevertheless, these results indicate that disturbance events not only initiate diversity of canopy-forming algae by thinning monospecific canopies, but through the liberation of available substratum.

Variation at local scales need not impede tests for broader scale patterns

Ecologists are not always mindful of the constraints imposed by their scale of observation and sometimes prematurely attempt broad generalisations or become mesmerised by local details depending on their predilections. We specifically chose a character that is known for its local and unpredictable variation (morphology of kelp) to test the effect of scale on our ability to determine spatial patterns. We compared the morphology of Ecklonia radiata between monospecific and mixed stands of canopy-forming algae across temperate Australia (>5,100 km coastal distance) within a hierarchy of several spatial scales. While E. radiata specimens were generally larger in monospecific than in mixed stands, we failed to observe differences in morphology between stands at many sites and locations. Despite substantial local variation, differences between stands became increasingly clear at broader scales. The frequency of inconsistent differences between stands was greatest at local scales (sites separated by kms), intermediate at intermediate scales (locations separated by 100s of kms) and least at regional scales (regions separated by 1,000s of kms). These observations support the idea that large scale patterns can emerge from apparent stochasticity at small scales, and that unaccountable variation at local scales need not impede tests for similar patterns at broader scales. Most ecologists work at scales where complexity tends to be greatest (i.e. local) and is likely to be explained by special and unique events. It is encouraging, therefore, to observe that patterns can emerge from complexity at local scales to provide new opportunities to answer some of the more interesting questions about the relative importance of processes across the vast parts of the world’s coast.

The prevalence and production of turf-forming algae on a temperate subtidal coast

M. Copertino, S.D. Connell and A. Cheshire. 2005. The prevalence and production of turf-forming algae on a temperate subtidal coast. Phycologia 44: 241–248.This study shows that canopy-forming algae, composed mainly of fucoids and the kelp Ecklonia radiata (both Phaeophycea), dominated space on South Australian coasts relative to turf-forming algae. However, where canopy-forming algae are absent turf-forming algae are the primary occupiers of space (~ 70%). On some reefs where canopy-forming algae are restricted in spatial extent, turfs can occupy as much as 40% of reefs. Turf-forming algae are an abundant component of algal assemblages, but relatively little is known about their contribution to the primary productivity on temperate reefs, relative to canopy-forming species. This study reveals that net productivity rates of turfs at one South Australian location were very high across depths (1.3–2.9 g C m−2 day−1 or 23.2–88.0 mg C g ash-free dry weight−1 day−1), comparable to the values discovered on tropical reefs. Although turf-forming algae are much more productive than canopy-forming algae on a biomass basis, the annual net production per area is two to seven times lower for turfs than for canopy-forming algae. However, if negligible exudation rates are assumed for turfs (up to 1%), the biomass of carbon produced by turf algae represents 44–71% of the carbon incorporated into biomass of kelps. Taken together, these results suggest that while canopy-forming algae can be correctly assumed to be the major source of total carbon produced on temperate reefs, the contribution of turf-forming algae may be substantial to the biomass production and turnover on South Australian reefs.

Persistence of the exotic kelp Undaria pinnatifida does not depend on sea urchin grazing

We investigated mechanisms enabling persistence of the introduced Asian kelp Undaria pinnatifida on the sea urchin (Heliocidaris erythrogramma) 'barrens' on the east coast of Tasmania. Development of dense stands of U. pinnatifida requires disturbance to reduce the cover of native algae. Observations of U. pinnatifida occurring abundantly on sea urchin barrens suggests that disturbance in the form of grazing by sea urchins prevents recovery of native canopy-forming species, allowing dense stands of U. pinnatifida to persist. We examined this hypothesis over a 30 mo period in a manipulative experiment in which the response of native algae and U. pinnatifida was examined in treatments comprising all possible combinations of +/- urchins, +/- U. pinnatifida and +/- enhanced native algal spore inoculum. The results demonstrated that the sea urchin H. erythrogramma can have a significant impact on U. pinnatifida abundance. The response was most dramatic in the 2001 sporophyte growth season, when sea urchins destructively grazed U. pinnatifida sporophytes in experimental plots on the urchin barren. In other years, when there was higher recruitment of U. pinnatifida sporophytes, urchins reduced sporophyte abundance but did not prevent development of a U. pinnatifida canopy. Removal of sea urchins resulted in a slow increase in cover of understorey red algae, but only limited recovery of native canopy-forming species. In treatments where both sea urchins and U. pinnatifida were removed, cover of canopy-forming species did not exceed 6% over the duration of the study. Thus, in the absence of sea urchin grazing, there was no evidence of inhibition of U. pinnatifida by native algae. While the intensity of sea urchin grazing may directly influence the extent of the U. pinnatifida canopy, recovery of native canopyforming species was apparently influenced by a combination of factors including seaurchin grazing, depth and, most importantly, the degree of sediment accumulation on the rocky substratum. The manipulations showed that removal of the factor (i.e. sea urchin grazing) that ostensibly facilitated replacement of native canopy-forming algae by U. pinnatifida did not realise recovery of native canopy-forming species.